Biological systems exhibit fluctuations driven by active (energy consuming) processes, such as molecular motors in the cytoskeleton and flagella propelling bacteria. It is tempting to treat these fluctuations in the same manner that thermal fluctuations are treated in equilibrium systems. Recent experiments have compared these inherent fluctuations with the response to small external perturbations. From these measurements, an effective temperature may be defined using the fluctuation-dissipation formalism of equilibrium statistical mechanics. In active systems, this effective temperature is frequency dependent (unlike the thermal case), and is higher than the ambient temperature. Another parameter that is useful for characterizing deviations from equilibrium of such fluctuations is the kurtosis of their distribution function; deviations of the kurtosis from its value for a Gaussian distribution indicate that the system may not be mapped onto an equilibrium ensemble.We address these issues theoretically by examining simple model systems of randomly kicked “particles”, with a variable number of kicking “motors”. Our generalized particles and motors represent different objects in different real systems, and we make specific choices when comparing to experimental observations. We exactly calculate the non-equilibrium fluctuation-dissipation relations and resulting effective temperatures. We show that their frequency dependence is model dependent and generally non-monotonous. We investigate the non-Gaussianity of the velocity distribution functions by a combination of numerical simulations and theoretical estimates. We show that the kurtosis has a non-monotonous dependence on the motor activity. We identify situations in which these two indicators of non-equilibrium behavior contradict. In particular, in the presence of multiple motors, the kurtosis of the distribution recovers its Gaussian value, while the effective temperature may still exhibit strong frequency dependence. We compare our results with two recent measurements of oscillations of the red-blood cell membrane, where these two indicators where independently measured.
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